INSECTA
MUNDI
A Journal of World Insect Systematics
0098
A new fossil species of stag beetle from Dominican Republic amber,
with Australasian connections (Coleoptera: Lucanidae)
Robert E. Woodruff
Florida State Collection of Arthropods
P.O. Box 147100
Gainesville, Florida 32614-7100
Date of Issue: September 25, 2009
CENTER FOR SYSTEMATIC ENTOMOLOGY, INC., Gainesville, FL
Robert E. Woodruff
A new fossil species of stag beetle from Dominican Republic amber,
with Australasian connections (Coleoptera: Lucanidae)
Insecta Mundi 0098: 1-10
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2009
0098: 1-10
A new fossil species of stag beetle from Dominican Republic amber,
with Australasian connections (Coleoptera: Lucanidae)
Robert E. Woodruff
Florida State Collection of Arthropods
P.O. Box 147100
Gainesville, Florida 32614-7100
Abstract. The first New World amber member of the family Lucanidae is described from the Dominican Republic. Its age is presumed to be Miocene (20-30 million YBP). It is also the fourth known amber species, the second
Miocene fossil species, the second fossil species in the subfamily Syndesinae, and the first species (fossil or
extant) of Lucanidae from the entire Caribbean. It is especially interesting because it is a member of the
Australasian genus Syndesus MacLeay. Other such disjunct Dominican amber insect fossils are also discussed.
Introduction
My first encounter with the specimen herein described was during a visit to the Dominican Republic
in 1983 to assist the late Jacob Brodzinsky in identifying his extensive Dominican amber collection. He
had set aside this fossil as an unknown beetle family. I immediately recognized it as a member of the
family Lucanidae and requested a loan for later study. He added it to my registry of Dominican amber
fossils as #5656.
Based on my initial identification, it was mentioned in a list by Poinar (1992: 285). Ratcliffe and
Ocampo (2001) indicated that I was in the process of describing this fossil, but nothing further has been
published. Subsequently I borrowed the specimen for description, but shortly thereafter the entire Brodzinsky
collection was purchased by the Smithsonian Institution (Davis 1989), and I was asked to return my loan.
The collection was originally housed in the Entomology Department of the U.S. National Museum, but
later transferred to the Department of Paleobiology, where it now resides.
During a study trip to the Smithsonian in 1998, I was able to examine the specimen again and to
compare it with extant members of the Lucanidae in the National Collection. Sometime between my first
examination (Fig. 3) and the current study, the piece was broken and repaired (Fig. 8). Certain features
were obscured by the fractures, making description more difficult. Not until the use of the Auto-Montage
became a reality, which greatly assists in illustrating amber inclusions, was I encouraged to complete
this description.
Fossil Lucanidae. The following summary is taken from a recent catalogue of fossil Scarabaeoidea
(Krell 2007: 2-5), which listed 17 species of fossil stag beetles. These are distributed in 14 genera in 5
subfamilies. The oldest are known from the Jurassic of Mongolia (Nikolajev 1990), 1 of which was included in a separate family, the Paralucanidae (Nikolajev 2007). Ceruchus fuchsii Wickham (1911) and a
single elytron of Lucanus fossilis Wickham (1913) are the only 2 previously known New World fossil
species. Both are compression (flattened) specimens from the Florissant, Colorado shales and considered
Oligocene in age. Wickham (1920, 1927, 1933) provided a catalogue and supplements to North American
fossil Coleoptera. Horning (undated) provided a website bibliography of the Lucanidae of the world.
Geologically (from oldest to most recent) and geographically, the fossil Lucanidae are here summarized: Upper Jurassic (Mongolia, 2 spp.); Lower Cretaceous (Russia, 3 spp.); Upper Cretaceous (Kazakhstan,
1 sp.); Eocene (Germany, Czechoslovakia, 5 spp.); Oligocene (Germany, Russia, Colorado, 5 spp.); Miocene
(France 1 sp.).
Only 3 species are listed as amber fossils, and all those were described from Baltic amber, listed as
Eocene in age (Motschulsky 1856, Waga 1883, Zang 1905). Ceruchus fuchsii Wickham is the only other
fossil member of the subfamily Syndesinae listed. Thus, of the family Lucanidae, S. ambericus n. sp.
constitutes the fourth known amber species, the first New World amber species, the second Miocene fossil
species, the second fossil species in the subfamily Syndesinae, and the first species (fossil or extant) from
the entire Caribbean.
1
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WOODRUFF
Syndesus ambericus Woodruff, new species
Holotype. Amber fossil, Dominican Republic, probably Cordillera Septentrional. Deposited in the
Brodzinsky-Lopez Peña Collection, USNM #502873, Acc. # 371429, Woodruff # 5656, housed in the Department of Paleobiology, National Museum of Natural History, Washington, D.C. Enclosed in the same
piece is a fly of the family Scatopsidae.
General Description. Specimen contained in amber, cut in cabochon form of elongate oval, convex on
one side (referred to as dorsal) and relatively flat on the reverse (ventral). Dimensions: 37.5mm long,
20.5mm wide, 7.1mm thick (all maximum measurements); beetle fossil 13mm long (from mandible tip to
abdomen tip), ca. 4mm high, width unmeasurable. Amber dark orange (darkened since first examined).
Fractures presently obscuring many features, but Fig. 3 shows the specimen as it appeared in 1984. Body
cylindrical, convex, elongate, with coarse punctures on pronotum and elytra. Habitus remarkably similar (compare Fig. 3 and 5, lateral view) to the extant S. cornutus (Fabricius).
Head broad, eyes large, prominent, nearly round. Antennal club large, with 7 lamellae (Fig. 7); other
antennomeres not visible. Head laterally with sharp projections in front of eyes (similar to Fig. 6). Maxillary palpi long, two-thirds mandible length, filiform, 3-segmented. Labial palpi 2-segmented, length half
that of maxillary palpi. Mandibles prominent, symmetrical, horn-like, with 3 teeth (Fig. 4, 12), length
two-thirds pronotal length.
Pronotum (Fig. 3, 10) convex, cylindrical, glabrous, coarsely punctate (most punctures deep and
contiguous). Marginal line evident, appearing continuous laterally and posteriorly (no dorsal view). Medially with small projection, not carinate or sharply delineated, but clearly visible; concave below to head.
Length two-thirds elytral length. Central furrow suggested in lateral view, but dorsal view obscured.
Elytra (Fig. 3, 8, 9) convex, cylindrical, glabrous; intervals (likely 10, difficult to count) prominently
raised, intervening punctures coarse, deep, nearly contiguous. Fractures presently distorting and obscuring view. Pygidium not visible, but fine golden setae projecting at elytral margin. Margin appearing
complete, terminating with smoothly rounded, gradual curve.
Legs similar to modern S. cornutus. Anterior tibiae dentate with 5 prominent teeth, becoming larger
anteriorly, small teeth interspersed between each; spur broad, terminal.
Sex. Based on the sexual dimorphism in the Syndesinae, and shape of the mandibles, the specimen
appears to be a male.
Comparisons. In most respects the fossil is so remarkably similar to the modern S. cornutus that there
can be little doubt about the generic placement. Obvious specific differences are found in the mandibles,
pronotal punctation, and pronotal projection (horn). The coarse, nearly contiguous pronotal punctures
easily distiguish it from any known species.The mandibles of S. cornutus (Fig.1, 5, 6) are more elongate
and with only 2 teeth (points), whereas S. ambericus (Fig. 4, 8, 12) mandibles are broader and have 3
teeth. The pronotal projection in S. cornutus (which is the origin of the specific name) is far more prominent and carinate (Fig.1, 5, 6) than in S. ambericus (Fig. 3). Few other character states are sufficiently
visible in the fossil to distinguish it.
Etymology. The commercial amber fossil company formed by Jacob and Marianella Brodzinsky (Woodruff 2004b) was incorporated as “Amberica”, combining “Amber” and “America”. I have chosen to honor
them for providing important fossils to the scientific community, by Latinizing their well-known company name.
Systematics. The family Lucanidae is not a large one by beetle standards. It has long been considered
the ancestral stock within the Scarabaeoidea, and it contains many species with relict distributions. The
latest checklist (Paulsen 2008), indicated 4 subfamilies, 109 genera, about 800 species, with 196 New
World species in 32 genera, with only 24 Nearctic species in 8 genera. Paulsen (2005) stated that “ The
classification of the family at the tribal level is currently superfluous...” There are no current keys to
species in many genera (including Syndesus MacLeay).
The genus Syndesus was created as the type of a new family by MacLeay (1819). In their treatment
of the Lucanidae of the World, Didier and Séguy (1952, 1953) considered it as a subfamily (Syndesinae)
A NEW FOSSIL STAG BEETLE FROM THE DOMINICAN REPUBLIC
1
INSECTA MUNDI 0098, September 2009 • 3
2
4
3
5
6
Figure 1-6. Syndesus spp. 1, 2, 5, 6: Syndesus cornutus (Fab.). 1) Left lateral view of head and pronotum (note
prominent mandibles and palpi). 2) Enlargement of eye and antenna (note 7 lamellae in club). 5) Habitus,
lateral. 6) Habitus, dorsal. 3-4: Syndesus ambericus Woodruff, n. sp. 3) Amber fossil; habitus lateral in 1983. 4)
Enlargement of right mandible (note 3 “teeth”).
4 • INSECTA MUNDI 0098, September 2009
WOODRUFF
with only 2 genera (Syndesus and Psilodon Perty). Within Syndesus, they recognized 5 species: 2 from
Australia, 2 New Caledonia, and 1 erroneously from Africa (Benesh 1955: 73); in Psilodon (with
Hexaphyllum Grey as a synonym) were 3 species from South America. In Psilodon, Martinez and ReyesCastillo (1985) and Boucher (1993) added new species from Brazil and Bolivia, respectively.
In his catalogue, Maes (1992) listed the subfamily Syndesinae to include Syndesus as the single
genus (with Psilodon and Hexaphyllum as synonyms) of a separate tribe (Syndesini), along with
Sinodendonini and Ceruchini. With this classification, the South American members (Psilodon) are combined with the Australasian ones, obscuring the disjunct distribution and morphological differences.
Paulsen (2008) stated that “A meaningful classification of the Lucanidae at the tribal level cannot be
created until all genera are properly diagnosed, and morphological and molecular phylogenetic analyses of
the higher-level lucanid relationships are conducted.” That process was begun by Paulsen and Hawks
(2008) when they described the new tribe Platyceroidini based on “...preliminary molecular
analysis...(unpublished)” and morphological differences with Platycerini.
Ratcliffe (2002: 8) listed Syndesus and Psilodon as distinct. In his website checklist of 2005, Paulsen
included the genus Psilodon as a synonym of Syndesus. However, in the most recent version (Paulsen
2008), the 2 genera are considered distinct. Several authors have weighed in on this classification (Holloway
1960, 1968; Howden and Lawrence 1974; Kikuta 1986; Krajcik 2001; Mizunuma and Nagai 1994; Smith
2006), but there seems to be little agreement. As recent as 1993, a new species (P. gilberti Boucher) was
described from Bolivia (Nor Yungas at 1500 meters). Unlike all other South American species of Psilodon,
it has 7 lamellae in the antennal club, showing its relationship to the Australasian Syndesus. In fact,
pronotal and mandible characters, illustrated in Boucher’s figures 6-7, show a remarkable similarity to
S. cornutus from Australia. A thorough revision of both “genera” will be required to clarify their status.
It is not my intent to evaluate classifications here, but to point out that the number of antennal lamellae
(6 in Psilodon, 7 in Syndesus) would appear to be a sufficient morphological character (along with their
distributions) to separate these 2 “genera”. Although I have not examined all the species in either genus,
Paulsen (2006) indicated that one species of Syndesus (S. cancellatus Montrouzier) has a 6-segmented
club.
Provenance. There can be little doubt that this fossil is from the Dominican Republic. Brodzinsky never
dealt in amber from elsewhere (Woodruff 2004b). Although he tried diligently to obtain specific locality
data from the miners and suppliers, it was usually futile or unreliable. Because of color, inclusions, and
vendor certainty, some pieces can be associated with specific mines. This specimen appears to have come
from the Cordillera Septentrional (not the eastern, younger deposits). Brodzinsky said (personal communication) that it likely came from the famous Palo Quemado mine, located north of Santiago, near La
Cumbre.
Age. There are no scientific techniques for dating amber itself. However, minimum ages can be obtained
from associated microfossils in the sedimentary strata in which the amber is found. In the Dominican
Republic, deposits were classified in the Altamira formation, once thought to be Eocene (Hueber and
Langenheim 1986). Lambert et al. (1985) and Poinar and Canatella (1987) reported on a frog, which was
dated by a controversial technique (MRI) as Eocene. However, most mines in the Cordillera Septentrional
are now considered to be marine sediments of Miocene age (Grimaldi 1995; Iturralde-Vinent and McPhee
1996; Poinar 1992; Woodruff 1994). However, Dilcher et al. (1992) suggested that “...the amber clasts,
from all physical characteristics, were already matured amber at the time of redeposition into marine
basins. Therefore, the age of the amber is greater than Miocene and quite likely is as early as late
Eocene”.
The age of Hispaniola continues to be controversial. Donnelly (1988) indicated that the Greater Antilles
started emerging 105 million years ago, with a rapid increase in size about 80 million years ago. Ross and
Scortese (1988) indicated that the Caribbean tectonic activity began in the Apian (118 million years ago),
and Hispaniola may have begun to break from Cuba during the Oligocene (35 million years ago). Their
exact size and position during subsequent geological periods is unknown, but most geologists agree that
the Caribbean plate has moved many times, and individual islands occupied very different positions from
today.
A NEW FOSSIL STAG BEETLE FROM THE DOMINICAN REPUBLIC
INSECTA MUNDI 0098, September 2009 • 5
7
8
10
9
11
12
Figure 7-11. Syndesus ambericus Woodruff, n. sp. 7) Antennal club (note 7 lamellae in club). 8) Lateral view,
habitus; note fractures in 2009. 9) Enlargement of elytral punctures, right side. 10) Enlargement of pronotum;
note dense, coarse punctures. 11) “Minute black scavenger fly” (Scatopsidae), located 5mm away from the Syndesus
ambericus holotype. 12) Right lateral view of head and pronotum; note mandibles and palpi.
6 • INSECTA MUNDI 0098, September 2009
WOODRUFF
As I indicated earlier (Woodruff In Woodruff and Sanderson 2004), Hispaniola is composed of 2 separate tectonic plates, the southern one subducting under the northern main island. On this southern plate
there is concrete evidence that land existed in the late Cretaceous (De León 1989) of Hispaniola. Fossil
trees are found in this deposit in the southern island that are replaced by the volcanic mineral pectolite
(var. Larimar; Woodruff 1986; Woodruff and Fritsch 1989). All amber deposits are found in the “north
island”, but no amber has been discovered on this “south island” or on the Haitian side of the border.
Biogeographical significance. When such a disjunct presence of a species of Lucanidae is discovered
as a fossil, and its relatives are geographically far removed, the initial question is “where did it originate?”
West Indian biogeography has been a significant area of research in recent times, involving many geologists and biologists (Woods 1989). However, many questions remain unresolved. The occurrence of various animal and plant species on each island have been documented both by dispersal and vicariance.
Many fossil species (and entire groups) are no longer present.
The amber insect fossils offer a unique clue to these origins, and continue to supply our oldest evidence of past Caribbean history. Sanderson and Farr (1960) were the first to call attention to the rich
amber fauna in the Dominican Republic, and the first Caribbean insect fossil was described a short time
later (Wille and Chandler 1964). A beautiful book, covering amber worldwide, was published by Grimaldi
(1996). In the first catalogue of the Dominican Arthropod fossils ((Perez-Gelabert 1999), 508 species in 301
genera are recorded. In his latest magnificent catalogue of the Hispaniolan fauna (Perez-Gelabert 2008),
he recorded 6,833 extant and (unbelievably) 1,404 fossil species.
Because I was aware that no member of the family Lucanidae was known from the Caribbean, I was
surprised to see this amber fossil from the Dominican Republic. When I later discovered its relationship
to the Australasian fauna, I was even more incredulous. However, recent studies of Dominican amber
fossils provide a growing list of similar distributions and relationships. Because they establish a pattern
of ancient extinctions and biogeographical evidence, special examples are summarized below.
1. Leptomyrmex neotropicus Baroni-Urbani (1980), and later (Baroni-Urbani and Saunders 1982). The
first fossil insect with Australian connections to be described from Dominican amber was this unusual
ant. The genus is presently confined to Australia, New Guinea, and New Caledonia, leading Wilson (1985)
to doubt the original placement of this amber fossil in the genus Leptomyrmex, although it was later
confirmed (Baroni-Urbani and Wilson 1987). Baroni-Urbani (1980) suggested that the disjunct distribution was the result of the genus having a “...former Tertiary cosmopolitanism or wide distribution...and a
post Miocene contraction of the area.”
2. Valeseguya disjuncta Grimaldi (1991). This woodgnat of the family Anisopodidae is a common amber
fossil in the Dominican Republic. In fact, what was originally believed to be this species (subsequently
identified as a species of Scatopsidae by D.A. Grimaldi 2009, in litt.) is present (Fig. 3, 11) in the amber
piece with the Lucanid fossil described here. The type of the genus and the single other species is based on
a unique extant male from Australia (Colless 1990). Such a gap in distribution is not easily understood,
except by one-time faunas being connected and subsequent extinctions. Grimaldi (1991: 13) emphasized
that “There is no doubt about the close relationship between the extinct (Dominican) and living (Australian) species.”
3. Mastotermes electrodominicus (Krishna and Grimaldi 1991). This primitive, giant termite was described from Dominican amber, and it has 1 living relative; from Australia and New Guinea (another
species has been described from Mexican amber). In Australia the living species thrives and is considered
a serious pest species. This fossil was one of the first amber pieces to have DNA extracted (DeSalle et al.
1992). Although retaining many roach-like features, the DNA studies suggested that these fossils were
true termites and not “missing links” with the roaches (Grimaldi 1996). In a recent paper (Inward et al.
2007), comprehensive molecular studies show that termites are eusocial cockroaches; they relegate the
Order Isoptera to a family (Termitidae) within the Order Blattodea.
4. Brachypsectra vivafosile (Woodruff 2002 [2004a]). When this unusual beetle was discovered, the family
Brachypsectridae contained only 3 known species, with relict distributions (India, Singapore, and South-
A NEW FOSSIL STAG BEETLE FROM THE DOMINICAN REPUBLIC
INSECTA MUNDI 0098, September 2009 • 7
western United States. Two larvae had been found in Dominican amber, but no adults. For this reason,
the extant adult specimens were therefore considered “living fossils” (i.e., vivafosile). An adult was found
subsequently in Dominican amber and described as Brachypsectra moronei Branham (2006). Recently a
modern larva of the family was discovered in Australia (Costa et al. 2006), but has not been named for
lack of adults. It does establish additional evidence for the Australian-Dominican connection.
Extinction. Such distributions appear to be explained only by invoking theories of continental drift and
plate tectonics. Grimaldi (1991) refers to this phenomenon as “geographical extinctions”. For the genus
Syndesus (and many of the insects mentioned above) to be present in the Miocene Caribbean and modern
Australasia, their ancestors must have once existed on the same land mass of the southern continent
Gondwanaland, after the first division of the supercontinent Pangaea. These insects are extremely ancient relics of a much broader distribution!
Curiously, this ancient stag beetle (and any other species of Lucanidae present in the Miocene) became extinct in the Caribbean, while other scarab beetles seemed to proliferate, speciate, and evolve into
island endemics. Presently, the scarab genus Phyllophaga contains 48 Hispaniolan species, all of which
are confined to that island (Woodruff and Sanderson 2004). Relationships to mainland faunas are obscure.
Although the genus has hundreds of species in Central America, each Caribbean island appears to have
mutually exclusive species.
Biology. Although there is no direct evidence for the behavior, biology, or habitats of this fossil, there is
considerable general interest in postulating prehistoric environments (Poinar 1999). This species probably lived in similar forests to extant species of the genus Syndesus. Most species of Lucanidae live in
decaying wood, and often in larger tree trunks. Most are presently rare, localized, and difficult to locate;
some (e.g., European stag beetle) are currently on endangered species lists. Their relict distributions
include most climatic zones, except the Arctic and Antarctic. Virtually none has ever been considered of
economic concern, although Lawrence (1981) reported an incident of Syndesus cornutus damaging a
bridge support in Canberra, Australia.
Monteith (in litt. 2009) provided the following details of the biology of S. cornutus in Australia and
Tasmania: It is one of the “really common species” in eastern Australia from Tasmania in the South to
tropical Queensland in the north. It is found mostly in rainforests, but also occurs in Eucalyptus forests,
especially at higher and damper sites. It breeds in wood with red rot fungi, mostly in large logs where big
colonies build up and may persist for many years in the same log. Adults are often found in the galleries
with the larvae. Adults come to lights “fairly regularly”.
Hueber and Langenhein (1986) pointed out that the tree which produced most Dominican amber was
related to African ancestors, and it was later described as a new species by Poinar (1991). Dilcher et al.
(1992) reported further on the leguminous trees from Dominican amber. The botanical inclusions are
numerous, and many await taxonomic study. It appears that both insects and plants on the island of
Hispaniola have some of their closest relatives currently living elsewhere!
Acknowledgments
I am indebted to the late Jake Brodzinsky for initially calling my attention to this specimen. I thank
the following National Museum personnel for their assistance: Terry Erwin, David Furth, and Garry
Hevel of the Entomology Department, and Jann Thompson and Conrad Labandeira of the Paleobiology
Department for their courtesy in making the specimen available for study. Paul Skelley assisted with the
Auto-Montage and the editorial process. Geoff Monteith provided valuable biological data on S. cornutus
in Australia. I thank Frank Krell, David Grimaldi, and M. J. Paulsen for their careful peer reviews. This
is Florida Department of Agriculture and Consumer Services, Division of Plant Industry, Entomology
Contribution No. 1158.
8 • INSECTA MUNDI 0098, September 2009
WOODRUFF
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Received August 17, 2009; Accepted September 11, 2009.